1 Vienna 20 1 A new planktonic foraminifera species ...cies Globigerina hagni, Globanomalina wilcoxensis globulosa and Globorotalia mattseensis were described by Gohrbandt (1967).

Abstract

The newly discovered planktonic foraminifer Hantkenina gohrbandti nov. spec. forms the evolutionary link between the genera

Clavigerinella and Hantkenina. This ancestor of the genus Hantkenina is characterized by pointed chamber ends with a nub (proto-

tubulospines) and in some cases by the first tubulospines appearing in a juvenile growth stage. The transition from Clavigerinella

caucasica to Hantkenina mexicana is observed in a 2 m thick part of the Middle Eocene Holzhäusl section (Mattsee, Austria) within

planktonic foraminifera Zone E8 and calcareous nannoplankton Sub-Zone NP15b. The section was deposited in bathyal water-

depths in the northwestern Tethyan realm and is now part of the Ultrahelvetic thrust unit.

Die neu entdeckte planktonische Foraminiferenart Hantkenina gohrbandti nov.spec. ist das evolutionäre Bindeglied zwischen den

Gattungen Clavigerinella und Hantkenina. Sie ist charakterisiert durch spitze Kammerenden mit einem distalen, hohlen Knoten

(Proto-Tubulospine) und in wenigen Fällen bereits mit einem echten Hohlstachel (Tubulospine) im juvenilen Wachstumsstadium.

Diese Evolution findet in einem nur 2m dicken Abschnitt des mittel-eozänen Holzhäusl Profils statt, innerhalb der Foraminiferenzone

E8, bzw. der kalkigen Nannoplankton Sub-Zone NP15b. Die bathyalen Sedimente des Holzhäusl-Grabens wurden in der nordwest-

lichen Tethys abgelagert und sind jetzt Teil des ultrahelvetischen Deckenkomplexes.

__________________________________

_____________________________________

KEYWORDS

Ultrahelvetic thrust unitplanktonic foraminifera

Northwestern TethysMiddle Eocene

Eastern AlpsHantkenina

1)*) 2)Fred RÖGL & Hans EGGER

1) Museum of Natural History, Burgring 7, A-1010 Vienna, Austria;

2) Geological Survey of Austria, Neulinggasse 38, A-1030 Vienna, Austria;

*) Corresponding author, [email protected]

1. Introduction

The planktonic foraminifer genus Hantkenina is characterized

by planispiral coiling and hollow chamber extensions, called

tubulospines. It evolved gradually from the genus Clavigeri-

nella, which shows radial elongate, clavate or digitate cham-

bers, but no tubulospines. This evolutionary trend and the

transition from Clavigerinella to Hantkenina was demonstrated

from the Austrian Holzhäusl section (Coxall et al., 2003) and

from the Kilwa drill sites in Tanzania (Pearson et al., 2004). At

both localities, a newly discovered species, which has been

named Hantkenina singanoae by Coxall and Pearson (2006),

was considered to be the missing link between the two genera.

The chambers of this species terminate in a distal hood (proto-

tubulospine), and it appears unclear how, and unlikely that,

straight tubulospines of the younger Hantkenina species could

evolve from this bent feature.

The material of the Holzhäusl section used by the English

working group is stored at the Natural History Museum in Vi-

enna and was sampled by K.H. Gohrbandt in the 1960s. The

precise position of his sample locations was unclear until H.

Egger re-sampled the site. Rögl and Egger (2010) report on

__________________________

the finding of a newly discovered

Hantkenina species with a first oc-

currence (FO) slightly below the FO

of H. singanoae. The stratigraphic

ranges of both species overlap. The

chambers of the Hantkenina nov.

spec. terminate in strongly pointed

tips or nubs, and in some rare juve-

nile specimens the first real tubu-

lospines have been observed. This

species is considered to be the real

ancestor of the genus Hantkenina.

Our paper gives the palaeontologi-

cal description of this important spe-

cies, for which the new name Hant-

kenina gohrbandti is introduced. The

holotype and paratypes are docu-

_

Austrian Journal of Earth Sciences ViennaVolume 104/1 2011

Figure 1: Location of the Holzhäusl section near Mattsee (Austria)._______________________

A new planktonic foraminifera species ( nov. spec.) from the Middle Eocene of the

northwestern Tethys (Mattsee, Austria)

Hantkeninagohrbandti

________________

Hantkeninagohrbandti

mented. The SEM stubs and thin-sections are stored in the

Micropalaeontological Collection of the Department of Geo-

logy & Palaeontology of the Museum of Natural History Vien-

na (inventory numbers 2011/0001 to 2011/0005).

The Holzhäusl section (47°58´26´´N; 13°07´09´´E) is located

near Mattsee, 20 km north of the town of Salzburg in Austria

(Fig. 1). It is part of the Ultrahelvetic thrust unit (UH) of the

Eastern Alps. The UH originates from the passive southern

margin of the European Plate and was deposited in bathyal

water depths at approximately 35° northern palaeolatitude in

the northwestern Tethyan realm. The Holzhäusl section is part

of the informal lithostratigraphic unit “Buntmergelserie” and

consists predominantly of grey marlstone displaying average

carbonate content of 58wt%. In previous studies of this loca-

lity, the first records of Clavigerinella and Hantkenina in the

Austrian-Bavarian Helvetikum were mentioned (Thalmann,

1951, Aberer and Braumüller, 1958, Hagn, 1960). From the

abundant planktonic foraminiferal assemblage the new spe-

cies Globigerina hagni, Globanomalina wilcoxensis globulosa

and Globorotalia mattseensis were described by Gohrbandt

(1967). The evolution of Hantkenina took place in a 2 m thick

part of the succession in the lower part of the course of the

Holzhäusl creek.

Based on the planktonic foraminifera assemblage with Gu-

embelitrioides nuttalli, Igorina broedermanni and Hantkenina

singanoae, the Holzhäusl section is assigned to Zone E8 in

the zonation scheme of Berggren and Pearson (2005) and

Wade et al. (2010). In the calcareous nannoplankton zonation

scheme of Martini (1971), and its modified version by Aubry

(1991), the outcrop can be assigned to Zone NP15, Sub-Zone

NP15b (Sullivania gigas Sub-Zone), due to the occurrences

of Nannotetrina fulgens and Sullivania gigas.

Order Foraminiferida Eichwald, 1830

Suborder Rotaliina Delage & Hérouard, 1896

Superfamily Globigerinaceae Carpenter, Parker & Jones, 1862

Family Hantkeninidae Cushman, 1927

Genus Hantkenina Cushman, 1924

Hantkenina gohrbandti Rögl & Egger, nov. spec.

Plate 1, Figs 1-7; Plate 2, Figs 1-6, 9; Plate 3, Figs 1-8; Plate

4, Figs 4-5

Planispirally

coiled, laterally compressed, bi-umbilicate, four and a half

chambers in the final whorl, rapidly increasing in size; first

chamber small and broken, second chamber rounded, third

chamber ovate, fourth and fifth chambers radially elongate

with pointed chamber ends, final chamber with a thickened

nub; aperture a high equatorial arch with broad lips.

Planispirally coiled, inner whorl with a slight

trochospiral tendency, laterally compressed, bi-umbilicate;

__________

____________________________________

_____________

_______

2. Setting of sample location

3. Stratigraphy

4. Systematic description

Description of holotype (pl. 1, fig. 1, pl. 3, fig. 1):

Test morphology:

peripheral outline stellate; 4½-6 chambers in the final whorl

(first and second chamber commonly crushed in our material);

chambers rounded in early stage, later ovate to triangular,

extend rapidly in radial direction; the final and commonly also

penultimate chamber end in a sharp tip or thickened nub. The

nub appears as a rounded, hollow and perforated structure

(pl. 4, fig. 5). Some juvenile specimens show the first occur-

rences of real tubulospines (pl. 2, figs 4-6, 9, pl. 3, figs 10-

11). Interestingly, after developing a tubulospine in the inner

whorl, the following chambers are pointed or with a thickened

nub. In thin-section the hollow tubulospine can be observed

(pl. 4, fig. 4). From the material of K.H. Gohrbandt primitive

tubulospines were described already by Coxall et al. (2003,

pl. 6, figs 9, 16). Later, the specimens were placed in the new

species H. singanoae (Coxall and Pearson, 2006, pl. 8.13,

figs 16-17).

Weakly cancellate and perforate, non-spinose,

pustules may be present (pl. 1, fig. 6, pl. 3, figs 6-8). In some

final chambers fine furrows towards the pointed end are pre-

sent (pl. 1, fig. 7, pl. 3, fig. 3).

holotype: maximum diameter 684 microns; paratypes

407 to 860 microns.

Named in honour of Klaus H. Gohrbandt (Gulf

Breeze, Florida, USA; former employee of Rohöl-Aufsuchungs

AG, Vienna) for his fundamental work on the Paleogene of the

Helvetikum north of Salzburg.

The new species Hantkenina gohrband-

ti is distinguished from Clavigerinella, especially C. caucasica

Subbotina by the development of pointed chamber ends with

a nub in the youngest chambers, forming a prototubulospine

in sense of Coxall and Pearson (2006). In contrast to H. sin-

ganoae Pearson & Coxall (in Coxall and Pearson, 2006), the

straight and pointed chamber ends differ clearly from the cy-

lindrical, commonly hood-like chamber ends in H. singanoae

(comp. pl. 3, fig. 13 of the holotype of H. singanoae). Primi-

tive forms of H. mexicana, originally described as H. nuttalli

Toumarkine, have broad triangular chambers, where the cham-

bers continue without break in the blunt tubulospines (pl. 3,

figs 14-15). Intermittent forms between H. gohrbandti and H.

mexicana (forma nuttalli) show blunt, somewhat irregular tu-

bulospines (pl. 2, figs 7-8). The transition between both spe-

cies is also visible in the small grooves along proto-tubulos-

pines and real tubulospines (pl. 3, figs 3, 11-12), which may

correspond to protoplasmatic structures.

In the evolution from Clavigerinella

to Hantkenina the new species is a transitional form between

C. caucasica and H. mexicana. In C. caucasica rounded cham-

ber ends are present throughout the final whorl (comp. pl. 4,

fig. 1). The new species H. gohrbandti shows initially rounded

chambers in the juvenile stage, followed by ovate-elongate

chambers, and finally chambers with a pointed tip, ending com-

monly in a hollow nub (comp. pl. 4, fig. 5). In some instances

a true tubulospine is already developed in juvenile chambers

(comp. pl. 4, fig. 4). In H. mexicana true tubulospines are con-

stantly developed in the later chambers of the final whorl.

________________________________________

_________________________

_________________________________

__________________________

_________________

___

Type of wall:

Size:

Etymology:

Distinguishing features:

Phylogenetic relationship:

Fred RÖGL & Hans EGGER

A new planktonic foraminifera species (Hantkenina gohrbandti nov. spec.) from the Middle Eocene of the northwestern Tethys (Mattsee, Austria)

Discussion:

Stratigraphic range:

Geographic distribution:

Acknowledgements

References

Aberer, F. and Braumüller, E., 1958.

Aubry, M.-P., 1991.

Berggren, W.A. and Pearson, P.N., 2005.

Coxall and Pearson (2006) discussed the position

of their new species H. singanoae in comparison to Clavige-

rinella. The development of proto-tubulospines and terminal

nubs is different in all species of Clavigerinella and forms a

transition to later species of Hantkenina. As demonstrated by

Rögl & Egger (2010) in the material from the Holzhäusl sec-

tion a more clear transition to H. mexicana can be explained

by the straight and pointed chamber ends in H. gohrbandti. In

the case of H. gohrbandti proto-tubulospines and terminal nubs

are developed, supporting its assignment to the genus Hant-

kenina. The species H. singanoae is also present in our sam-

ples, but in small numbers (pl. 1, fig. 9), and the characteristic

hood is rudimentarily developed (pl. 3, fig. 9).

Middle Eocene, planktonic foraminifera

Zone E8 (Guembelitrioides nuttalli Lowest-occurrence Zone),

Nannotetrina fulgens Zone NP15, Sullivania gigas Subzone

NP15b. New calibrations of Zone E8 with the younger part of

magneto-chron C21n and C20r yield an estimated age span

of 46.4 to 44.4 Ma (Wade et al., 2011).

At present, this species is known only

from a short interval in the Holzhäusl section, township Matt-

see N of Salzburg, Austria, belonging to the Ultrahelvetic thrust

unit of the Eastern Alps.

Firstly, we have to thank Klaus H. Gohrbandt (Gulf Breeze,

Florida, USA) for providing us with his samples from the Holz-

häusl section. For discussions on the evolution and systematic

position of the transitional forms between Clavigerinella and

Hantkenina we are grateful to Helen Coxall and Paul Pearson

(Cardiff University, UK), to whom we have to extend our gra-

titude for providing us with SEM figures of the holotype and

unpublished material from Tanzania. We thank Helga Prie-

walder and Sabine Giesswein for their help in taking the SEM

pictures and Oleg Mandic for his assistance in taking pictures

in the light microscope. We are indebted to William A. Berg-

gren (Rutgers University, New Jersey, USA) and Ewa Malata

(Jagiellonian University, Kraków, Poland) for the constructive

reviews.

Über Helvetikum und Flysch

im Raume nördlich Salzburg. Mitteilungen der Geologischen

Gesellschaft in Wien, 49 (1956), 1-39.

Sequence stratigraphy: Eustasy or tectonic

imprint?. Journal of Geophysical Research, 96, 6641-6679.

A revised tropical to

subtropical Paleogene planktonic foraminiferal zonation. Jour-

nal of Foraminiferal Research, 35, 279-298.

_____________

__________________

______________________________

___________________

__

______________

Coxall, H.K., Huber, B.T. and Pearson, P.N., 2003.

Coxall, H.K. and Pearson, P.N., 2006.

Gohrbandt, K.H.A., 1967.

Hagn, H., 1960.

Martini, E., 1971.

Pearson, P.N., Nicholas, C.J., Singano, J.M., Bown, P.R., Co-

xall, H.K., Dongen van, B.E., Huber, B.T., Karega, A., Lees,

J.A., Msaky, E., Pancost, R.D., Pearson, M., and Roberts, A.P.,

2004.

Rögl, F. and Egger, H., 2010.

Thalmann, H.E., 1951.

Wade, B.S., Pearson, P.N., Berggren, W.A. and Pälike, H.,

2010.

Origin and

morphology of the Eocene planktonic foraminifer Hantkenina.

Journal of Foraminiferal Research, 33, 237-261.

Taxonomy, biostratigra-

phy, and phylogeny of the Hantkeninidae (Clavigerinella, Han-

tkenina, and Cribrohantkenina). Cushman Foundation Special

Publication, 41, 213-256.

Some new planktonic foraminiferal

species from the Austrian Eocene. Micropaleontology, 13, 319-

326.

Die stratigraphischen, paläogeographischen

und tektonischen Beziehungen zwischen Molasse und Helveti-

kum im östlichen Oberbayern. Geologica Bavarica, 44, 1-208.

Standard Tertiary and Quaternary calcare-

ous nannoplankton zonation. In: Farinacci, A. (Ed.), Procee-

dings II Planktonic Conference. Technoscienza, Roma, 739785.

Paleogene and Creatceous sediment cores from the

Kilwa and Lindi areas of coastal Tanzania: Tanzania Drilling

Project Sites 1-5. Journal African Earth Sciences, 39, 25-62.

The missing link in the evolutio-

nary origin of the foraminiferal genus Hantkenina and the pro-

blem of the Lower/Middle Eocene boundary. Geology, 38, 23-

26.

Mitteilungen über Foraminiferen IX. 44.

Vorkommen der Gattung Hastigerinella Cushman, 1927, im

österreichischen Mitteleozän. Eclogae Geologicae Helvetiae,

43, 223-224.

Review and revision of Cenozoic tropical planktonic fo-

raminiferal biostratigraphy and calibration to the geomagnetic

polarity and astronomical time scale. Earth-Science Reviews.,

doi: 10.1016/j.earscirev.2010.09.003.

___________

_____________________________

_

____________________

Received: 21 January 2011

Accepted: 1 April 2011

1)*) 2)Fred RÖGL & Hans EGGER1)

2)

*)

Museum of Natural History, Burgring 7, A-1010 Vienna, Austria;

Geological Survey of Austria, Neulinggasse 38, A-1030 Vienna, Aus-

tria;

Corresponding author, [email protected]

____

_______________



Plate 1:

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

All figured specimens are from the Mattsee-Holzhäusl section. Scale bar for all figures

200 microns.

Hantkenina gohrbandti nov. spec., holotype, lateral view, sample B5-08,

inv. no. 2011/0004/0011; maximum diameter 684 microns. Detail of final

chamber in pl. 3, fig. 1.

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample A2h-08,



Hantkenina gohrbandti nov. spec., paratype, apertural view, sample A2d-

08, inv. no. 2011/0004/0013; maximum hight 679 microns.

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample A2a-08,



Hantkenina gohrbandti nov. spec., paratype, lateral view, sample Go 64/1-

36/0, inv. no. 2011/0003/0006; maximum diameter 600 microns. The spe-

cimen was figured in COXALL et al. (2003, pl. 6, fig. 8) as Calvigerinella-

Hantkenina transition.

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample B4-08,


chamber in pl. 3, fig. 6; wall surface of earlier chamber in pl. 3, figs 7-8.

Hantkenina gohrbandti nov. spec., single chamber with distinct nub, sam-

ple A2h-08, inv. no. 2011/0004/0007; maximum hight 415 microns.

Clavigerinella caucasica (Subbotina), sample B2-08, inv. no. 2011/0002/

0014. Characteristic elongate, clavate chambers, rapidly increasing in hight.

Hantkenina singanoae Pearson & Coxall, sample B3-08, inv. no. 2011/0001/

0003. Final chamber ending with a hood (proto-tubulospine). Detail of final


_________________________________________

_________________________________________

_____________

_________________________________________

__________________________________________

_

_____

_________________________________________

1:

2:

3:

4:

5:

6:

7:

8:

9:



Plate 2:

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

All figured specimens are from the Mattsee-Holzhäusl section. Scale bar for all figures

200 microns.

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample A2e-08,

inv. no. 2011/0004/0002; maximum diameter 707 microns.

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample A2f-08,

inv. no. 2011/0004/0005; maximum diameter 860 microns.

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample B4-08, inv.

no. 2011/0001/0020; maximum diameter 654 microns. Detail of final cham-

ber in pl. 3, fig. 2.


no. 2011/0001/0014; maximum diameter 624 microns. First chamber of the

final whorl with real tubulospine, second chamber with a poined chamber

end, further chambers with broken tips (see pl. 3, fig. 10).


no. 2011/0001/0019; maximum diameter 408 microns. Juvenile specimen

with a tubulospine in the first chamber of final whorl; further chambers with

thickened conical knobs (see pl. 3, fig. 11).

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample A2f-08, inv.

no. 2011/0002/0002; maximum diameter 636 microns. Final chamber with

thickened knob, earlier chambers with pointed ends.

Hantkenina gohrbandti – H. mexicana transition, lateral view, sample Go

64/1-36/4b, inv. no. 2011/0003/0024; maximum diameter 880 microns. The

blunt tubulospine in the final whorl forms a transition between the proto-

tubulospine in H. gohrbandti and the slender tubulospine in Hantkenina

mexicana.

Hantkenina gohrbandti – H. mexicana transition, lateral view, sample Go

64/1-36/4b, inv. no. 2011/0003/0027; maximum diameter 690 microns. The

specimen was figured in COXALL et al. (2003, pl. 6, fig. 10, 17) as "primi-

tive" Hantkenina nuttalli. The tubulospine is somewhat deformed compa-

rable to that in fig. 7.

Hantkenina gohrbandti nov. spec., paratype, lateral view, sample Go 64/1-

36/4b, inv. no. 2011/0003/0021; maximum diameter 408 microns. The spe-

cimen was figured in Coxall et al. (2003, pl. 6, fig. 9, 16) as "primitive" Han-

tkenina nuttalli. The blunt proto-tubulospine forms a transition to that in H.

cf. mexicana. Pores are developed all along the process. Later, Coxall &

Pearson (2006, pl. 8.13, figs 16-17) placed it in H. singanoae.

_____________

_____________

______________________________________________

_____________

_________________________

_________________

____________________________________________________

___________________________________________

__________

1:

2:

3:

4:

5:

6:

7:

8:

9:



Plate 3:

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Hantkenina gohrbandti nov. spec., paratype, final chamber of specimen in

pl. 1, fig. 1.


pl. 2, fig. 3.


pl. 1, fig. 7. The pointed end shows fine radial grooves, similar to those

along the tubulospines of H. mexicana (comp. fig. 12).


pl. 1, fig. 4.


pl. 1, fig. 2.


pl. 1, fig. 6.

Hantkenina gohrbandti nov. spec., paratype, first complete chamber in the

final whorl of specimen in pl. 1, fig. 6, showing the development of small

pustules.

Hantkenina gohrbandti nov. spec., paratype, higher magnification of the wall

in fig. 7, specimen in pl. 1, fig. 6.

Hantkenina singanoae Pearson & Coxall, final chamber with a hood, spe-

cimen in pl. 1, fig. 9.

Hantkenina gohrbandti nov. spec., paratype, first and second chamber in

the final whorl of specimen in pl. 2, fig. 4.

Hantkenina gohrbandti nov. spec., paratype, conical knob at the end of the

prelast chamber of specimen in pl. 2, fig. 5.

Hantkenina mexicana Cushman, tubulospine of the final chamber in speci-

men pl. 3, fig. 15, showing fine radial grooves (protoplasmatic structures?)

and pores up to the compact concial end.

Hantkenina singanoae Pearson & Coxall, 2006, holotype. SEM figure kindly

provided by Paul Pearson.

Hantkenina cf. mexicana Cushman "forma H. nuttalli Toumarkine", with

broad chambers in the stellate arrangement of H. mexicana; sample A1-

08, inv. no. 2011/0001/0012.

Hantkenina mexicana Cushman; sample A2f-08, inv. no. 2011/0002/0006.

Primitive form with elongated chambers, continuing in stout tubulospines

(comp. fig. 12).

________________

_________________________________

__________________________

_________________________

__________________________

______________________________________

_____________________________________

1:

2:

3:

4:

5:

6:

7:

8:

9:

10:

11:

12:

13:

14:

15:



Plate 4:

Figure

Figure

Figure

Figure

Clavigerinella caucasica (Subbotina), thin-section, sample B4-08, inv. no.

2011/0005/0001. In thin-section the chamber ends are rounded, without a

thickening or a nub (maximum diameter of the specimen: 0.43 mm).

Hantkenina mexicana Cushman, thin-section, sample Rö 2-98, inv. no.

2011/0005/0002. The slender chambers end in a hollow tubulospine, in

the inner whorl small tubulospines can be observed already (maximum

diameter of the specimen: 0.72 mm).

Hantkenina gohrbandti nov. spec., thin-section of a juvenile specimen with

a tubulospine and a nub (arrow); sample B4-08, inv. no. 2011/0005/0003.

Maximum diameter of the specimen: 0.31 mm.

Hantkenina gohrbandti nov. spec., thin-section. Pointed chamber with

nearly circular nub (arrow); sample A2e-08, inv. no 2011/0005/0004. Ma-

ximum diameter of the specimen: 0.44 mm.

____

____________________________

_____________________

_______________________

1:

2-3:

4:

5:

1 Vienna 20 1 A new planktonic foraminifera species ...cies Globigerina hagni, Globanomalina wilcoxensis globulosa and Globorotalia mattseensis were described by Gohrbandt (1967).

Documents